ECO~ORJ of Freshwuttr Fish 1997: 6: 3 6 4 4 Copyright Q Munksgaard 1997

Printed in Denmark . All rights reserved

ECOLOGY OF
FRESHWATER FISH
ISSN 0906-GG91

Lateral migration of fishes in Amazon

floodplains
Fernandes CC. Lateral migration of fishes in Amazon floodplains. C. C. Fernandes
Ecology of Freshwater Fish 1997: 6: 3 6 4 4 . 0Munksgaard, 1997 lnstituto Nacional de Pesquisas da Arnazdnia
(INPA), Manaus, Brazil
Abstract - Migrations of fish in the outlet channel of Lago do Rei, a flood-
plain lake on Careiro Island in the Amazon River, were examined using three
capture methods. Of the 85 species found in the Parana do Rei (channel), 20
migrated between the lake and river. Migrations from the lake to the river oc-
curred in two pulses. In August-September, as the river water level began to
recede, some fish migrated to the river with the current. As the river water
level rose in December-January, additional fish migrated towards the main
river against the current. Fish returned to the lake by March or April after
they had spawned. The migratory patterns of six species, family Curimatidae Key words: migration; Amazon River:
and Prochilodontidae: Potamorhina latior, Psectrogaster amazonica, PI ruti- Curirnatidae; Prochilodontidae
loides, Curimata kneri, Prochilodus nigricans and Sernaprochiladus taeniu- Cristina Cox Fernandes, lnstituto Nacional de
rus are described in detail. Lateral migrations of fish in this ecosystem appear Pesquisas da Amazsnia, Alarneda Cosme Ferreira
to be closely linked to breeding and to physical habitat changes related to 1756 Aleixo 69083,Manaus AM, Brazil
river water level. Accepted for publication March 7, 1996

Un resumen en espaiiol se incluye detrhs del texto principal de este articulo.

Many fishes undergo seasonal migrations. In gen-
eral, migrations include longitudinal movements,
along the main body of rivers, and lateral migra-
tions, between a river and its floodplain (Welcomme
1985). Many authors have reported longitudinal mi-
gration in characiform fishes of the seasonal flood-
plains of the Amazon Basin (Godoy 1979; Goulding
1980; Worthman 1982; Ribeiro 1983; Zaniboni
1985; Bayley & Petrere 1989; Menezes & Vazzoler
1992). However, lateral migrations in the Amazon
have rarely been studied (Goulding 1980; Ribeiro
1983). Daget (1958) and Welcomme (1985) charac-
terized such movements for fishes of seasonally
flooded African rivers. Here I present the first study
of lateral migrations in the Amazon.
Along the Amazon River, the seasonal water level
fluctuation is quite large, with a mean variation of
approximately 10 m per year (Junk 1983). Flood
pulse is driven primarily by seasonal precipitation
cycles in the upstream catchment rather than by lo-
cal rainfall and is the major force regulating biota in
river-floodplain systems (Junk 1984; Junk, Bayley
& Sparks 1989). There is, however, little documen-
tation of specific movements of animals in response
to the seasonal flux of water level. Many fishes from
floodplains exhibit morphological, physiological
and biochemical adaptations that permit their sur-
vival during periods of drought and anoxic water
(Junk et al. 1983; Saint-Paul 1984). Migratory be-
haviors may be a means to avoid such temporary ad-
verse conditions and they have been related to re-
production and the utilization of allochthonous food
resources (Goulding 1980).
The objective of this study was to document and
discuss fish lateral migration patterns in a channel
connecting a floodplain lake to the Amazon River. I
specifically address (1 ) the fish assemblage compo-
sition of the study area: (2) patterns of lateral move-
ments of these fishes; and (3) detailed patterns of
migration of 6 species of Curamatidae and Prochilo-
dontidae.
Methods
Migration patterns and their physiological correlates
were assessed using (1) net captures; (2) direct ob-
servations of fish movements; (3) observations of
reproductive chronology; (4) catch records from

36

subsistence and commercial fisheries, and ( 5 ) asso-
ciations of migration behavior with water level.
Study area
This work was carried out on Careiro Island (Stern-
berg 1956; Annibal 1983; Anonymous 1988), the
first island downstream from the confluence of the
Rios Solim6es and Negro (Fig. 1). The island is sit-
uated at 3" 05' to 3" 12' S and 59" 35' to 59" 50' W
and has an area of 470 km2.Its greatest length is 41
km and greatest width is 17 km. The island is bor-
dered on the north by the Rio Amazonas and by the
Parana do Careiro on the south.
Careiro Island is a typical example of a season-
ally flooded, fluvial plain (viirzea), composed of al-
luvial sediments (Guillaumet et al. 1988). Its flat to-
pography is marked by low levees along the edges
surrounding shallow lakes and natural channels
(paranas) connecting lakes and river.
The largest lake on Careiro Island, Lago do Rei,
is permanent and has a surface area of about 100
km2 during high water (Fig. 1). The central part of
the lake is permanent and the marginal area sur-
rounding the lake consists of seasonally flooded for-
est and fields. The main river determines the regime
of waters in the floodplain lakes, invading the lakes
during the rainy season and draining its waters dur-
ing the dry season (Bayley & Petrere 1989). Parana
do Rei (Fig. 1) is the largest channel connecting the
Rio Amazonas and the Lago do Rei system. It is ap-
proximately 20 km long, 35 m mean wide, and its
depth is 2-12 m, varying with the hydrologic period
(Petry 1989). The direction and control of the flow
in the Parana do Rei alternates with variation of the
Fig. 1. Map of Careiro island, showing location of Parana do
Rei, Lago do Rei and Rio Amazonas (dry season).
Lateral migration of fishes in Amazon floodplains
n I
30/2500
"1 2000
5
3
1500 ,E
-
VI
m
3
lo00 2-u
-
E
500
0
j a s o n i j fmamj j a s o n d j fmamj
1985 1986 , 1987
Fig. 2. Water level from July 1985 through June 1987 (line)
based on daily records for the Rio Negro, provided by the Ad-
ministration of the Port of Manaus (PORTOBRAS), and total
monthly number of individuals of at least 22 species of subsist-
ence fishery on the Parana do Rei (histogram).
levels of the Lago do Rei and Rio Amazonas. Dur-
ing the beginning of the fall to the dry season, the
current in the Parana do Rei moves from the lake to-
wards the river. At this time of the year, the parana
is the only connection between the Amazon and
Lago do Rei. During the high water, the current
pulses from the Amazon to the Lago do Rei, and
Parana do Rei has the Amazon river characteristics
and discharges 750 million m3 in the lake (Anony-
mous 1988). During this period, the Amazon River
overflows above the low edges of Careiro island (on
about 25 m height) and the lake becomes connected
to the river through different areas. Parana do Rei is
also connected through IgarapC do Irau-ACu to the
small Lago Joari, but this lake is not connected to
the Amazon, except during the floods.
Fig. 2 illustrates the water level flux during 1985
and 1986 based on daily records for the Rio Negro
provided by the Administration of the Port of
Manaus, Brazil (Portobras), located less than 60 km
up river from the island. The current in the channel
carries a large quantity of organic material (tree
trunks, branches and floating islands of grass). Gen-
erally, from the end of the floods until the beginning
of the dry season (October-November), access to the
lake through the channel is interrupted by a dense
layer of aquatic macrophytes (Eichorniu crussipes
and Echinochloa polystuchya) and woody matter. At
this time, the water of the parana stagnates and the
deficit of oxygen is high because of decomposition of
vegetation from the land, submersed during the
floods. This barrier (about 5 km long) is broken by the
force of the flow as water comes out of the lake, and
37
Fernandes
can kill fish as they move through the channel (see The common cast net was fished alone for 1 h, or
also Annibal1983; Odinetz-Collart & Moreira 1989). both cast nets were used for 30 min each. Fish catch
Odinetz-Collart & Moreira (1989) characterized per unit of effort (CPUE) among nets was standard-
the seasonal limnology of Parana do Rei among ized to number of individuals per hour per 100 m2 of
other places in Careiro Island, in 1987. The maxi- gill net and to number of individuals per hour of cast
mum oxygen concentration is found in January, June nets.
and October (9-12.1 mg.l-'), while the minimum The direction in which fishes were migrating, in
occurs in March and at the end of August to the be-
ginning of September (1-3 mg - I-'). Daily variations
of the dissolved oxygen show that there is an in-
crease during the day, with the maximum between
2-8 pm, at the surface and at 2 m depth. Conductiv-
ity ranged between 46 pS-cm-'. 1-' in September and
79 yS-cm-'.I-' in December, and pH varied be-
tween 5.2 (July-August) and 7.1 (March). No signif-
icant seasonal variation was found in surface water
temperature (26.7-29.4"C).

Experimental procedure
Tagging methods, which are often used for the study
of migration in temperate fishes, have proven unsuc-
a s o n d j f m a m j j a s
cessful in studies of neotropical fishes (Godoy 1979;
Worthman 1982; Carvalho 1983). Petrere (1985)
suggested that the low recapture rates of Amazon
River fishes is related to high mortality and disper-
sion of fish across a broad area. In this study, migra-
tion patterns were assessed using net captures and
direct observations.

Gill and cast net sampling

Four sizes of drifting gill nets were used: (1) 40 mm
stretch mesh, 17.20 m long, and 2.70 m diameter; (2)
60 mm stretch mesh, 17.20 m long and 2.25 m diam-
eter; ( 3 ) 80 mm stretch mesh, 20.00 m long and 3.00 a s o n d j f m a m j j a s
m diameter; (4) 110 mm stretch mesh, 32.50 m long
and 2.35 m diameter. Gill nets werc stretched across
the water body, maintained by two canoes, and al- c)
lowed to drift with the direction of the current. Cast m 40-
nets were used to sample along the channel margins 3-
.-E
ring cast net
where aquatic plants were abundant. Common cast 30 -
nets (25 mm mesh and 3.0 m diameter) were used to
fish on the surface on all the field trips, and a "ring"
cast net (25 mm mesh and 3.0 m diameter) that closes 20 -
at depth was used only during the flooded period.
Fish were sampled in the eastern half of the channel
(Fig. I), every other week from August 1985 to Sep- 10 -
tember 1986. The distance over which samples were
taken varied depending on the direction and the ve- .=
locity of the current. In August 1985 only one type of a s o n d j f m a m j j a s
gill net (mesh size 60 mm) was used. In September 1985 1986
1985, as waters began to recede, conditions made the Fig. 3. Total catch per unit of effort (CPUE) of 87 species
use of gill nets impossible, then only cast net sampling summed for 2-week periods in outlet channel of Lago do Rei us-
was used. ing (a) gill nets (* in September 1985 no gill nets were used),
Each sample comprised four 15-min gill net sets. (b) common cast nets, and ( c ) ring cast nets.

38
 Lateral migration of fishes in Amazon floodplains
December (exiting or entering Lago do Rei), was @ Potornorhim bfiw @ Pwctropmtrr rwtiloidre

determined from the side of gill nets on which fish

were captured. Each gill net sample was considered
an independent replicate and is represented in terms
of numbers of individuals per species per month.
Direct observations also gave information about the
direction of movement, as conditions allowed (e.g.,
Harden Jones 1968). In general, movements of
characiforms are near the surface to mid-depth
(Goulding 1979; Bonetto et al. 1981; Ribeiro 1983;
Zaniboni 1985). In spite of the turbidity of the wa-
ter, I was able to see individuals of the migratory
species reported here moving near the surface, iden-
tify them and determine the direction of their migra-
tion. 1985 1986 1885 1986

Samples of specimens of each of six focus spe-

cies, Potamorhina latior, Psectrogaster amazon-
ica, Psectrogaster rutiloides, Curimata kneri,
Prochilodus nigricans and Semaprochilodus taeni-
urus, were assessed for sex and maturity. This
was done because sex and maturity might be a cor-
relate or cause of migration. Sexual maturity was
estimated, in the field, after dissection of the ab-
dominal cavity, according to the following (Nikol-
sky 1963): (0) Immature - gonads small, thin and
transparent; (1) Non-mature - gonads thicker, but
still poorly developed, sex recognition difficult to
determine; (2) Maturing - gonads enlarged, testes
white, ovaries orange but ovules not yet visible to
the naked eye; (3) Mature - ovules opaque, visi-
ble to the naked eye and testes enlarged; (4)
Prespawning - ovules translucent, milt flows
when pressure applied; ( 5 ) Spent or post-spawn-
ing - ovaries long, usually folded but occupying a
large portion of the abdominal cavity, only scat-
tered remains of reproductive cells, testes soft and
empty. Relative frequencies were calculated for the
six stages of maturity to obtain an indication of the
reproductive status of the population.
Voucher specimens of all species were kept for
identification purposes in the fish collection at
INPA.

Subsistence and commercial fishery data

Daily catch of subsistence fisheries was collected
for the Parana do Rei through interviews with fish-
ers (Anonymous 1988) from July 1985 through June
1987. I determined the total number of individuals
collected per month during this period.
Regional commercial fishery data have been col-
lected since 1976 by the Division of Fisheries Re-
sources of INPA, in the Municipal Market of
Manaus. Fishers in the channel only use seines to
catch migratory species, but this gear is not used in
the Lago do Rei because of tree trunks and
Fig. 4. Monthly relative frequency of the maturity stages for fe-
males and males of a) Potamorhina latior, b) Psectrogaster
rutiloides, c) Psectrogaster amazonica, d ) Curimata kneri, e )
Semaprochilodus taeniurus and f) Prochilodus nigricans. Hori-
zontal bars refer to immature gonads, circles to non-mature,
white bars to maturing, diagonal hatching to prespawning and
vertical lines to spent or post-spawning gonads.
branches. Therefore, only the seine data (biomass,
kg) from the channel were used as an additional
source of information on the presence of fish.

39
Fernandes
Table 1. Species present in the catch of the three gears in Parano do Rei. Numbers between parenthesis refer to the species family: (1) Osteoglossidae; (2)
Clupeidae; (3) Engraulidae; (4) Characidae; (5) Cynodontidae; (6) Hemiodontidae; (7) Erythrinidae; (8) Prochilodontidae; (9) Curimatidae; (10) Anostomidae; (11)
Apteronotidae; (12) Sternopygidae; (13) Rhamphichthyidae; (14) Doradidae; (15) Auchenipteridae; (16) Ageneiosidae; (17) Pimelodidae; (18) Hypophthalmidae;
(19) Cetopsidae; (20) Loricariidae; (21) Cichlidae; (22) Scianidae.

Gill Common Ring N Relative

Species nets cast net cast net total abundance

Triportheusalbus (4) 288 194 53 535 14.87

Mylossoma aureum (4) 284 197 3 484 13.45
Pofamorhina lafior (9) 239 199 20 458 12.73
Mylossoma duriventris (4) a6 252 0 338 9.39
Triportheusangulafus (4) 78 177 0 255 7.09
Pimelodus blochii (7) 1 223 1 225 6.25
Eigenmannina melanopogon (6) 171 50 0 221 6.1 4
Schizodon fasciatum/Leporinus frifasciafus (10) 16 100 2 iia 3.28
Pellona flavipinnis (2) 72 3 0 75 2.08
Psectrogaster rutiloides (9) 6 54 2 62 1.72
Ageneiosus ucayalensis (1 6) 56 4 0 60 1.67
Semaprochilodus taeniurus (a) 45 13 0 58 1.61
Prochilodus nigricans (8) ia 36 0 54 1.50
Curimafa kneri (9) 25 12 2 38 1.06
Pellona castelnaeana (2) 34 2 2 38 1.06
Pterygoptichthys multiradiatus (20) 1 33 3 37 1.03
Roeboides myersi (4) 10 21 1 32 0.89
Rhaphiodon vu$inus (5) 22 10 0 32 0.89
Triportheuselongalus (4) 24 6 0 30 0.83
Auchenipterus nuchalis (15) 19 7 0 26 0.72
Ageneiosus sp. (16) 24 0 1 25 0.69
Rhytiodus microlepis (10 ) 3 19 2 24 0.68
Sorubim lima (17) 1 21 0 22 0.61
Centromochlus heckeli (15) 18 1 1 20 0.55
Cynodon gibbus (4) 2 19 0 21 0.58
Hypophthalmus fimbriafus (18) 12 6 0 18 0.50
Hypophthalmus marginatus (I a) 17 0 0 17 0.47
Cetopsis caecutiens (19) 17 0 0 17 0.47
Hypophthalmus edentatus (18) 2 14 1 17 0.47
Centrodorasbrachyafus (14) 3 11 0 14 0.39
Potamorbifla altamazonica (9) 6 a 0 14 0.39
Serrasalmus rhombeus (4) 0 12 1 13 0.36
Osteoglossum bicirrhosum (1) 6 6 0 12 0.33
Tetragonopterus argenteus (4) 0 11 0 9 0.25
Brachyplatystoma vaillanfii (17) 2 6 0 a 0.22
Rhytiodus argenteofuscus (10 ) 1 7 0 8 0.22
Ageneiosus brevifilis (I a) 6 2 0 a 0.22
Plagioscion squamosissimus (22) 4 3 1 a 0.22
Pimelodus alfissimus (17) 0 4 4 a 0.22
Lycengraulis batesi (3) a 0 0 a 0.22
Pseudodoras niger (14) 0 7 0 7 0.19
Curimatella meyeri (9) 4 3 0 7 0.19
Hypostomus plecosfomus (20) 0 6 1 7 0.19
Pseudoplatystomafasciafum (17) 1 5 0 6 0.17
figenmannia sp. (12) 4 2 0 6 0.17
Pygocentrus nattereri (4) 0 5 0 5 0.14
Hemiodus microlepis (6) 5 0 0 5 0.14
Leporinus frederici (10 ) 0 5 0 5 0.14
Opsodoras humeralis (14) 4 1 0 5 0.14
Brycon eryfbropferum (4) 2 2 0 4 0.14
Anodus sp. (6) 1 0 3 4 0.14
Pterodoras granulosus (14) 4 0 0 4 0.14
Rhamphichthys marmoratus (13) 0 4 0 4 0.14
Parauchenipterus galeatus (1 5) 3 1 0 4 0.14
Chaefobranchopsis sp. (1 2) 2 1 0 4 0.14
Hoplerythrinus unifaeniatus (7) 4 0 0 4 0.14
28 other species 14 30 5 45 1.25

40

Results
I captured 3,646 individuals of 87 species represent-
ing 22 families (Table 1). Of these, 20 species were
clearly laterally migrating. An additional 10 species
were probably also migrating, but they were not ob-
served in sufficient numbers to be classified as mi-
gratory.
Migrations from the lake appeared to peak twice,
as measured by CPUE (Fig. 3). The first movement
occurred during the receding of the waters from Au-
gust to September 1985, when the currents in the
parana were from the lake to the main river. The
second occurred from December 1985 to January
1986, when current of the water in the parana was
from the Amazon to the lake, at the beginning of the
floods. Of the six focus species (Table 2), Potamor-
phina latior, Psectrogaster amazonica, Curimata
kneri exhibited periods of increased abundance in
August-September 1985 and December 1985,
Semaprochilodus taeniurus and Prochilodus nigri-
cans yielded low catches in August to September
1985 and peaks of capture in December 1985; and
Psectrogaster rutiloides showed the reverse with a
high yield in August and total absence in December.
Between August 1985 and January 1986, fish
were observed schooling and migrating, from the
lake to the Amazon up-river. During December all
individuals were captured on the lake side of the gill
nets; for instance, during the second week of De-
cember, 249 individuals of the six focus species
were captured leaving the lake, while no individuals
of these species were found to be entering the lake.
In January and February some dispersed non-
schooling individuals of migratory species were ob-
served in gill net captures to be moving inward from
the river to the channel.
Lateral migration of fishes in Amazon floodplains
l? Latior; l? rutiloides, P. amazonica and C. kneri
showed advanced stages of maturation (mature and
prespawning) from November 1985 to April 1986
(Fig. 4). In both September 1985 and 1986, none of
these fishes exhibited ripe gonads, and several had
spent gonads. S. taeniurus and I! nigricans appeared
ready to reproduce in December 1985 and January
1986. In September and October of 1985, most indi-
viduals were spent or immature, although a few
were in maturing and prespawning stages.
Subsistence and commercial fishery data
The subsistence fishery yielded its highest capture
between September and April of both years (Fig. 2).
Each year the catch was lower between May and
August. Two peaks of capture occurred, with the
pattern of abundance similar to the gill net and cast
net data. The highest capture rates were during the
receding water in September 1985 and January
1986. These data corresponded to my own captures
(compare Fig. 2 and Fig. 3).
Fig. 5 illustrates commercial fishing capture in
the channel over a 10-year period, and also displays
the mean water level of the Rio Negro at Manaus. A
comparison of fish catch and water level suggests
that the catches are associated with the falling wa-
ter/dry season period, and also with the migration of
fish from the lake.
Discussion
This study suggests the following three-phase pat-
tern of lateral migration in the channel of Lago do
Rei.
( 1 ) In August - September during the recession of
waters, some groups of P. latior; 19 amazonica, l?

Table 2. Total capture per unit of effort (number of individuals per hour per 100 m2 of gill nets and number of individuals per hour of cast nets) for 6 focus species
using two fishing gears, between August 1985 and September 1986. Note that gill nets were not used in September 1985.

Months

Species Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep

Gill nets:
Potamorphina latior 588 - 0 0 743 22 3 7 0 0 0 0 0 0
Psectrogaster amazonica 0 - 0 0 36 3 0 3 0 3 0 0 0 0
Psectrogaster rutiloides 40 - 0 0 0 0 2 0 0 3 0 0 0 0
Curimata kneri 20 - 0 0 109 0 0 0 0 0 0 0 0 0
Prochilodus nigricans 0 - 0 0 42 0 0 0 0 0 0 0 0 0
Semaprochilodustaeniurus 0 - 0 0 174 21 0 0 0 0 0 0 0 0
Cast nets:
Potamorhina latior 6 32 0 2 25 25 14 9 6 0 0 0 0 8
Psectrogaster amazonica 1 10 0 0 1 1 0 1 4 0 0 0 0 1
Psectrogaster rutiloides 3 4 0 0 0 3 1 0 3 0 1 0 0 3
Curimata kneri 1 1 0 0 0 0 0 0 5 0 0 0 0 0
Prochilodus nigricans 0 3 4 0 1 0 0 0 0 0 0 0 0 8
Semaprochilodustaeniurus 1 1 0 0 0 0 0 0 0 0 0 0 0 2

41
Fernandes
"EAR
Fig. 5. Biweekly seine net catch in the Parana do Rei, related
with the level of the waters of the Rio Negro between 1976 and
1986. Water level from 1976 through 1986 based on daily
records for the Rio Negro, provided by the Administration of
the Port of Manaus (PORTOBRAS). Solid line refers to total
fish catch; dashed line to water level.
rutiloides, C. kneri, F! nigricans, and S. taeniums
leave the Lago do Rei swimming in the direction of
the Amazon River. The movement follows the same
direction of the currents of the channel. When the
fish reach the river, they go up river against the cur-
rent in the direction of the Rio Solimbes.
(2) At the beginning of rising water (December -
January), a second lateral migration occurs, appar-
ently related to reproduction, when the fish again
leave the floodplain lake and then migrate up the
river. These fishes are mature, and from this it is in-
ferred that their migration is related to spawning.
Since water in the channel is now flowing into the
lake, this migration involves swimming against the
current.
(3) After spawning (January-May), a lateral mi-
gration entering the recently flooded plains occurs,
in a dispersed way, without formation of schools.
The currents of the river favor larval drift and swim-
ming of spent adults through the floodplain which
presumably functions as feeding and nursery
grounds.
The first movement from the lake (August-Sep-
tember) does not appear to be related to reproduc-
tion. The majority of individuals captured at this
time had non-mature or spent gonads, but some indi-
viduals were in the initial stages of maturation, and
all had accumulations of fat in the abdominal cavity.
Once in the river these fishes initiate a longitudinal
migration up-river (Fernandes, personal observa-
tion). Goulding (1983) noted that the receding-water
period was the most productive for fishing on the
floodplains. The same was observed in this work.
The main families caught in the channel entrance at
that time were: Curimatidae, Characidae, Anostomi-
dae, Hemiodontidae and Pimelodidae (Table 1j.
Although the hydrological conditions in 1985-86
42
were 2 m above the long-term average during the
low water and high water, these does not seemed to
have interfered with the migration behavior. Fish
seem to migrate every year, in different concentra-
tions, in spite of the intensity of the flood pulse (Fig.
5).
Many authors have observed migrations from
tributaries at the onset of receding water. Migration
behavior might be initiated by changing food re-
sources (Shulman 1974; Saldaiia & Venables 1983).
For example, some fishes have been shown to feed
for many months in the inundated forest before
their migration (Goulding 1980; Goulding & Car-
valho 1982; Lowe-McConnell 1987). Maximum ca-
loric content of Prochilodus mariae in the Orinoco
river occurs at the beginning of the migratory sea-
son (Saldaiia & Venables 1983), and in general
maximum fat content of whole fish may reach 16 to
30% during high water (Junk 1984). Young
Sernaprochilodus, abandoning floodplain lakes dur-
ing receding waters and entering the Rio Negro,
have been shown to be fattened from their feeding
in floodplain lakes (Ribeiro 1983).
It is also possible that an oxygen gradient might
also serve as a stimulus for fish to leave the lake.
Odinetz-Collart & Moreira (1989) observed a drop
in the level of dissolved oxygen in the Lago do Rei,
during August - September 1987. I also measured
low levels of oxygen during the falling of water, in
the channel and near the entrance of the lake. The
water of the Parana do Rei was low in oxygen, pre-
sumably because of the large quantities of organic
material and detritus from the flooded terrestrial
vegetation available for decomposition. In addition,
dense aquatic plants and the inundated forest had re-
duced the insulation in the surface of the water dur-
ing flooding, also decreasing the production of oxy-
gen by the phytoplankton (Junk 1984). It is well
known that fish are capable of behavioral responses
to the oxygen concentration of water (Smith 1985).
In 1986, I observed a large number of moribund in-
dividuals of migratory species, moving from the
lake. Apparently these fish suffered from anoxia fol-
lowing an algae bloom, because they were breathing
in the superficial layer of the water. In general, sea-
sonal changes in depth, current speed, current direc-
tion, turbulence, conductivity, dissolved salt and ox-
ygen concentration may also serve as migratory
stimuli to fish.
The lateral migration exhibited by part of the fish
assemblage leaving the system favors the survival of
the remaining component. Because of the decrease
of the population density, the resources available to
the fish remaining will be greater. Some individuals
of these species might stay in the lake because they
are not in a suitable physiological condition to mi-

grate (e.g., insufficient fat reserves). Their chances
of survival in the lake during the dry period may be
improved because of the reduction of fish density.
Reproductive migration, e.g., the second pulse of
migration here, have been observed at the beginning
of floods by many researchers (Ribeiro 1983; Junk
1984; Carvalho & Merona 1986). Fish that initially
remained in the lake, caught from December to
March, were found to exhibit reproductive maturity.
This is consistent with the idea that when this sec-
ond pulse of fish reach the river, their longitudinal
upstream migration is reproductive in nature.
It is likely that spent individuals migrate from the
river into the floodplain and the forests to rebuild
their energy storage. As inferred from the low
CPUE, fish do not school as they re-enter the chan-
nel. Goulding (1980) and the fishers from the region
of the Rio Madeira also observed that spent fish do
not move in large schools, but move to the tributar-
ies in dispersed groups. Schools of Colossoma
macropomurn disperse after spawning and begin en-
tering the floodplain forest (Goulding & Carvalho
1982). Bonetto et al. (1981) found that the speed of
upstream reproductive migration of Pruchilodus
platensis and SaEminus maxilosus has to be fast, but
the movements back to the original area are slow,
with deviations for feeding or rest.
In conclusion, many authors agree that migration
is an adaptive behavior, related to seasonal habitat
selection and allowing for the use of optimal habi-
tats for development, growth and reproduction
(Northcote 1984; Lowe-McConnell 1987), and this
finding appears to apply to lateral migrations as
well. Thc benefits of migration in general should ex-
ceed its costs (energetic, risk of predation, or death
during movement) for such behavior to persist in a
species. This study also suggests that the costs and
benefits of lateral migrations might be more easily
detected than they can be for longitudinal migra-
tions, given the relative proximity and predictability
of seasonal habitats.
Resumen
1. He examinado la migracidn lateral de 10s peces en un canal
del Lago do Rei, situado en la Isla Careiro, en el Rio Amazonas
central (Brasil). Con este fin, se utilizaron tres mCtodos de cap-
tura diferentes. De las 85 especies encontradas en el Parana do
Rei (canal), 20 migraron entre el lago y el no.
2. Estas migraciones entre el lago y el rio se producen en dos
pulsos diferentes. En agosto-septiembre, cuando el nivel de
agua empieza a disminuir, algunos peces llegan hasta el n o ar-
rastrados por la coniente. A1 aumentar el nivel de agua en di
ciembre-enero, se produce una nueva migraci6n hacia el no,
contra la coniente. Los peces regresan a1 lago en marzo-abril,
despuCs de la freza.
3. En este articulo se describen en detalle 10s patrones migrato-
Lateral migration of fishes in Amazon floodplains
rios de seis especies de las familias Curimatidae y Prochilodon-
tidae: Potamorhina latior, Psectrogaster amazonica, I? ruti-
loides, Curimata kneri, Prochilodus nigricans y Semaprochilo-
dus taeniurus.
4. Parece que las migraciones laterales en este ecosistema esdn
asociadas a la reproduccidn y 10s cambios fisicos en el hhbitat
que se producen como consecuencia de 10s cambios en el nivel
del agua.
Acknowledgements
I gratefully acknowledge the endless patience of Jeff Podos and
John Lundberg in revising the manuscript, as well as input from
two anonymous reviewers. This work was part of a CNPq (Con-
selho Nacional de Pesquisa e Tecnologia), INPA (Instituto Na-
cional de Pesquisas da AmazBnia) and ORSTON (Institut
FranGais de Recherches pour le Development en Cooperation)
project, and was supported by the European Community (CEE).
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